Affinage

SMC4

Structural maintenance of chromosomes protein 4 · UniProt Q9NTJ3

Round 2 corrected
Length
1288 aa
Mass
147.2 kDa
Annotated
2026-04-28
64 papers in source corpus 15 papers cited in narrative 17 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SMC4 is a core ATPase subunit of the condensin I and condensin II complexes, where it heterodimerizes with SMC2 through its hinge domain and undergoes ATP-hydrolysis-coupled conformational changes that drive mitotic chromosome condensation and sister chromatid resolution (PMID:14532007, PMID:26904946, PMID:11267866). SMC4 nuclear accumulation is gated by CDK1 phosphorylation during mitosis, and its abundance is cell-cycle-regulated through APC/C-dependent proteasomal degradation at mitotic exit (PMID:10485849, PMID:30719224). Beyond its canonical condensin role, SMC4 functions as a transcriptional co-activator — for example at the PGAM1 and NEMO promoters — linking it to glycolytic reprogramming, NF-κB-dependent innate immunity, and, in cancer contexts, immune evasion through suppression of cGAS-STING signaling and upregulation of PD-L1 (PMID:37543034, PMID:29803706, PMID:40976049). SMC4 transcription is directly activated by FoxO1 and NFIA, and its mRNA is stabilized by METTL14-mediated m6A methylation (PMID:38403332, PMID:40933894).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 1999 High

    Establishing how CDK phosphorylation controls condensin function: Cdc2 directly phosphorylates SMC4 at T19, and this event is necessary and sufficient for mitosis-specific nuclear accumulation and chromosome condensation, linking cell-cycle kinase signaling to condensin activation.

    Evidence In vitro Cdc2 kinase assays, GFP localization of wild-type vs. T19A mutant Cut3/SMC4 in fission yeast

    PMID:10485849

    Open questions at the time
    • Phosphorylation sites in metazoan SMC4 not mapped in this study
    • Whether additional kinases contribute to SMC4 regulation in mitosis remains unresolved
  2. 2000 High

    Defining the human condensin complex: demonstration that human SMC4 (hCAP-C) and SMC2 form a heterodimer that associates with CNAP1/CAP-D2, establishing the existence of a condensin complex in human cells with mitosis-specific chromosome binding.

    Evidence Reciprocal co-immunoprecipitation from HeLa cell extracts, immunofluorescence localization through the cell cycle

    PMID:10958694

    Open questions at the time
    • Non-SMC subunit composition not fully delineated at this point
    • Whether multiple condensin complexes exist in vertebrates was unknown
  3. 2001 High

    Resolving whether SMC4 drives chromosome compaction versus sister chromatid resolution: Drosophila SMC4 loss-of-function reveals that SMC4 is specifically required for sister chromatid resolution, not axial shortening, distinguishing separable condensation functions.

    Evidence gluon mutant alleles in Drosophila with cytological analysis of mitotic chromosome structure

    PMID:11267866

    Open questions at the time
    • Whether this resolution function is separable from condensin I versus II was not addressed
    • Mechanism by which SMC4 promotes resolution at the molecular level not determined
  4. 2003 High

    Demonstrating that two distinct condensin complexes share the SMC2/SMC4 core: vertebrate cells contain condensin I and condensin II with different non-SMC subunits, different chromosome distributions, and distinct functional contributions to chromosome architecture.

    Evidence siRNA depletion in HeLa cells and Xenopus egg extract reconstitution with immunofluorescence

    PMID:14532007

    Open questions at the time
    • How the shared SMC2/SMC4 core is partitioned between the two complexes is unclear
    • Relative contributions to interphase versus mitotic functions not resolved
  5. 2015 Medium

    Structural insight into condensin architecture: cross-linking mass spectrometry revealed that the SMC2/SMC4 coiled coils can be closely apposed along their length both in vitro and in situ, and identified histones H2A and H4 as potential chromatin contacts.

    Evidence Amino acid-selective cross-linking MS with homology modelling on chicken SMC2/SMC4, including in situ cross-linking on mitotic chromosomes

    PMID:25716199

    Open questions at the time
    • Histone contacts not validated by orthogonal binding assays
    • Static cross-linking cannot capture dynamic conformational changes
  6. 2016 High

    Revealing the dynamic flexibility of the SMC2-SMC4 coiled coils: high-speed AFM showed the arms behave as highly flexible polymers that interconvert between multiple architectures, with head-to-hinge contacts, fundamentally revising the rigid-arm model of condensin.

    Evidence High-speed atomic force microscopy in liquid with quantitative polymer physics analysis of S. cerevisiae condensin

    PMID:26904946

    Open questions at the time
    • How flexibility relates to DNA loop extrusion activity was not tested
    • Nucleotide-state dependence of conformational dynamics not fully resolved in this study
  7. 2018 Medium

    Identifying a non-canonical transcriptional co-activator role for SMC4: in macrophages, SMC4 recruits H4K5 acetyltransferase activity to the NEMO promoter, epigenetically enhancing NF-κB and IRF3 activation and innate immune cytokine production.

    Evidence Epigenetic modifier screen, ChIP for H4K5ac at the nemo promoter, Smc4 knockdown in macrophages, in vivo sepsis model

    PMID:29803706

    Open questions at the time
    • Identity of the recruited acetyltransferase is unknown
    • Whether this function requires the intact condensin complex or monomeric SMC4 is unresolved
    • Not independently replicated
  8. 2019 Medium

    Establishing cell-cycle regulation of SMC4 protein abundance: yeast SMC4 is degraded at mitotic exit via APC/C and the proteasome, and unexpectedly the spindle checkpoint component Mad2 is required for this degradation, linking condensin turnover to checkpoint signaling.

    Evidence Cell cycle synchronization with APC/C and Mad2 genetic/chemical perturbation and proteasome inhibition in budding yeast

    PMID:30719224

    Open questions at the time
    • APC/C degron motif in SMC4 not mapped
    • Whether this mechanism is conserved in metazoans is untested
  9. 2022 Medium

    Linking SMC4 to pyroptotic cell death: TREM-1 directly interacts with SMC4 in septic cardiomyocytes, and this interaction promotes NLRP3 inflammasome activation and Gasdermin-D cleavage, placing SMC4 in a non-canonical inflammatory signaling pathway.

    Evidence Co-immunoprecipitation of TREM-1/SMC4, siRNA knockdown, LPS/nigericin pyroptosis model, in vivo CLP sepsis model

    PMID:36181338

    Open questions at the time
    • Reciprocal Co-IP direction not specified; binding domain mapping absent
    • Mechanism by which SMC4 activates NLRP3 inflammasome is undefined
  10. 2023 Medium

    Revealing how SMC4 attenuation reprograms cancer metabolism: reduced SMC4 in colorectal cancer upregulates glycolysis enzymes and lactate, driving histone lactylation that upregulates ABC transporters and chemoresistance; SMC4 also co-activates PGAM1 transcription, and combined SMC4/PGAM1 loss impairs F-actin assembly and causes cytokinesis failure.

    Evidence SMC4 knockdown with metabolic assays, histone lactylation ChIP, co-activation assays, F-actin imaging, polyploidy quantification in CRC cells

    PMID:37543034

    Open questions at the time
    • Whether SMC4 co-activation of PGAM1 requires condensin complex partners is unknown
    • Histone lactylation-ABC transporter link not independently confirmed
  11. 2024 Medium

    Defining upstream transcriptional and epitranscriptomic regulation of SMC4: FoxO1 binds the SMC4 promoter directly and also activates METTL14, which installs m6A marks on the SMC4 coding sequence to stabilize its mRNA, creating a coherent feed-forward loop controlling SMC4 expression.

    Evidence ChIP-seq and dual-luciferase for FoxO1-SMC4 promoter binding, METTL14 knockdown, m6A methylation mapping

    PMID:38403332

    Open questions at the time
    • m6A reader protein responsible for SMC4 mRNA stabilization not identified
    • Tissue specificity of the FoxO1-METTL14-SMC4 axis not determined
  12. 2025 Medium

    Expanding SMC4's oncogenic mechanisms: SMC4 suppresses cGAS-STING signaling to enable immune evasion in triple-negative breast cancer, upregulates PD-L1 in a STING-independent manner, interacts with GLUT1 to support glycolysis in prostate cancer via Rheb/mTOR, and is transcriptionally driven by NFIA to activate TGF-β/SMAD-mediated metastasis and LDHA-dependent aerobic glycolysis.

    Evidence SMC4 knockdown/overexpression with cGAS-STING and PD-L1 assays in 4T1 models; IP-MS identifying GLUT1 interaction validated by co-IP; NFIA ChIP and dual-luciferase at the SMC4 promoter with xenograft and metabolic assays

    PMID:40278414 PMID:40933894 PMID:40976049

    Open questions at the time
    • Whether immune evasion and metabolic functions depend on condensin complex integrity is unstudied
    • Direct versus indirect nature of SMC4-cGAS-STING suppression not distinguished
    • All cancer studies are from single labs and await independent replication

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unknown how SMC4's canonical condensin-dependent functions in chromosome condensation relate mechanistically to its emerging non-canonical roles in transcriptional co-activation, metabolic regulation, and immune signaling — specifically whether these require intact condensin complexes, monomeric SMC4, or distinct post-translational modification states.
  • No reconstitution assay distinguishing condensin-dependent from condensin-independent SMC4 activities
  • No structure of a non-canonical SMC4 complex
  • Physiological relevance of non-canonical functions outside cancer cell lines not established

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 3 GO:0140657 ATP-dependent activity 3 GO:0005198 structural molecule activity 2 GO:0140110 transcription regulator activity 2
Localization
GO:0005694 chromosome 5 GO:0005634 nucleus 3 GO:0005829 cytosol 1
Pathway
R-HSA-1640170 Cell Cycle 4 R-HSA-4839726 Chromatin organization 4 R-HSA-1430728 Metabolism 3 R-HSA-168256 Immune System 3
Complex memberships
condensin Icondensin II

Evidence

Reading pass · 17 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1999 Fission yeast condensin contains five subunits including Cut3/SMC4 and Cut14/SMC2, and Cdc2 kinase directly phosphorylates Cut3/SMC4 at T19 in vitro and in vivo (maximal at metaphase). Alanine substitution of T19 prevents nuclear accumulation during mitosis and causes condensation defects, demonstrating that Cdc2 phosphorylation of SMC4 is required for its mitosis-specific nuclear localization and chromosome condensation. In vitro Cdc2 kinase assay, in vivo phosphorylation analysis, GFP localization, temperature-sensitive mutant rescue, alanine substitution mutagenesis Genes & development High 10485849
2001 Drosophila SMC4 (encoded by the gluon locus) is required specifically for sister chromatid resolution during mitosis. Loss-of-function mutations allow normal longitudinal axis shortening but cause striking failure of sister chromatid resolution prior to anaphase, leading to chromosome breakage and apoptosis. DmSMC4 localizes to the axial chromosome core at metaphase/anaphase and dissociates during telophase decondensation. Genetic loss-of-function (gluon mutant alleles), immunolocalization, cytological analysis of mitotic chromosome structure Current biology : CB High 11267866
2000 Human SMC4 (hCAP-C) forms a heterodimeric complex with hCAP-E/SMC2 in HeLa cells. Co-immunoprecipitation of this complex identified CNAP1 (homolog of Xenopus XCAP-D2) as an interacting partner, establishing the existence of a condensin complex in human cells. Chromosome association of the hCAP-C-hCAP-E complex is mitosis-specific, with the complex sequestered in the cytoplasm during interphase, though a subpopulation remains on chromosomes as foci that colocalize with phospho-H3 clusters at late G2/early prophase. Co-immunoprecipitation from HeLa extracts, immunofluorescence localization, cell cycle analysis Molecular and cellular biology High 10958694
2003 Vertebrate cells contain two distinct condensin complexes (condensin I and condensin II) that share the same SMC2/SMC4 heterodimeric core but differ in their non-SMC subunits. siRNA depletion of condensin I- or condensin II-specific subunits produces distinct chromosome morphology defects; the two complexes show different distributions along the mitotic chromosome axis in vivo and in Xenopus egg extracts. siRNA depletion in HeLa cells, Xenopus egg extract reconstitution, immunofluorescence, biochemical fractionation Cell High 14532007
2015 Cross-linking mass spectrometry combined with molecular modelling of the chicken SMC2/SMC4 subcomplex reveals that the coiled-coil segments can be closely apposed along their lengths in isolated condensin complexes and in situ in mitotic chromosomes. Both histone H2A and H4 were identified as potential condensin chromatin-interaction partners via cross-linking data. Amino acid-selective cross-linking and mass spectrometry, homology modelling, in situ cross-linking on mitotic chromosomes Open biology Medium 25716199
2016 High-speed atomic force microscopy in liquid demonstrates that the Smc2-Smc4 coiled-coil arms of S. cerevisiae condensin are highly flexible polymers (persistence length ~4 nm) that dynamically interconvert between multiple architectures. The SMC head domains engage not only with each other but also with the hinge domain at the other end of the ~45-nm coiled coil. High-speed atomic force microscopy (AFM) in liquid, quantitative polymer physics analysis Cell reports High 26904946
2018 Macrophage Smc4 promotes inflammatory innate immune responses by epigenetically enhancing NEMO (IKKγ) transcription: Smc4 recruits H4K5 acetyltransferase activity to the nemo promoter, increasing H4K5 acetylation and enabling stronger NF-κB and IRF3 activation upon TLR or viral stimulation. Smc4 knockdown or deficiency reduced IL-6, TNF-α, and IFN-β production and protected mice from sepsis. High-throughput epigenetic modifier screen, ChIP for H4K5ac at nemo promoter, Smc4 knockdown in macrophages, in vivo sepsis model Journal of autoimmunity Medium 29803706
2019 Budding yeast Smc4 protein abundance is cell cycle regulated: it peaks at mitosis and falls to low levels in interphase. Smc4 degradation at the end of mitosis is dependent on the Anaphase Promoting Complex/Cyclosome (APC/C) and mediated by the proteasome. Unexpectedly, the Mad2 spindle checkpoint protein is required for mitotic Smc4 degradation. Overproduction of Smc4 delays decondensation but cannot promote premature condensation in interphase. Cell cycle synchronization, protein stability assays, APC/C and Mad2 genetic/chemical perturbation, proteasome inhibition Oncotarget Medium 30719224
2020 In Plasmodium spp., SMC2/SMC4 localize to centromeres (validated by co-localization with NDC80 and ChIP-seq) during early schizogony without forming condensin I or II complexes. In mature schizonts and during male gametogenesis, SMC2/SMC4 distribute diffusely and assemble into both condensin I and II complexes. Knockdown of smc2 and smc4 reveals essential roles in parasite proliferation and transmission. ChIP-seq, fluorescence microscopy co-localization with NDC80, conditional knockdown, parasite transmission assays Cell reports Medium 32049018
2023 SMC4 attenuation in colorectal cancer cells drives a diapause-like state by: (1) promoting expression of three investment-phase glycolysis enzymes and increasing lactate, which via histone lactylation upregulates ABC transporters causing chemoresistance; and (2) acting as a co-activator of PGAM1 transcription, such that combined SMC4/PGAM1 loss impairs F-actin assembly, causes cytokinesis failure and polyploidy, and inhibits proliferation. SMC4 knockdown/attenuation, metabolic assays (lactate, glycolysis), histone lactylation ChIP, transcriptional co-activation assays, F-actin imaging, polyploidy quantification Cell metabolism Medium 37543034
2022 TREM-1 interacts directly with SMC4 in cardiomyocytes under septic conditions, and inhibition of either TREM-1 or SMC4 prevents NLRP3 inflammasome activation and reduces Gasdermin-D cleavage, IL-1β, and caspase-1 cleavage, identifying SMC4 as a component of the TREM-1/NLRP3 pyroptosis pathway in cardiomyocytes. Co-immunoprecipitation (TREM-1/SMC4 interaction), siRNA knockdown, LPS/nigericin pyroptosis model, in vivo CLP sepsis model The FEBS journal Medium 36181338
2025 SMC4 promotes prostate cancer cell proliferation and metastasis through the Rheb/mTOR signaling pathway. IP-MS identifies GLUT1 (SLC2A1) as an SMC4 interactor, confirmed by co-IP, and SMC4 knockdown inhibits glycolysis rate and ATP production, linking SMC4 to metabolic regulation via GLUT1 interaction. CRISPR/Cas9 SMC4 knockdown, IP-MS interactome, co-IP validation of SMC4-GLUT1 interaction, glycolytic rate assay, in vivo murine lung metastasis model, mTOR/Rheb pathway western blotting Advanced science Medium 40278414
2025 SMC4 in triple-negative breast cancer maintains genomic stability to suppress cGAS-STING-mediated type I interferon production and CD8+ T cell recruitment; it also upregulates PD-L1 in a STING-independent manner. Knockdown of SMC4 restored tumor immunogenicity and potentiated anti-PD-1 therapy in immunocompetent 4T1 murine models. SMC4 knockdown/overexpression, cGAS-STING pathway assays, ELISA for cytokines, flow cytometry of CD8+ T cells, molecular docking, immunoprecipitation, in vivo 4T1 tumor model with anti-PD-1 International immunopharmacology Medium 40976049
2024 FoxO1 promotes SMC4 transcription by binding to the SMC4 promoter at the -1400/-1390 bp region (validated by ChIP-seq and dual-luciferase assay), and also transcriptionally activates METTL14, which increases SMC4 mRNA m6A methylation on its coding sequence, thereby increasing SMC4 mRNA abundance. ChIP-seq, dual-luciferase reporter assay, METTL14 knockdown, m6A methylation mapping, in vitro and in vivo proliferation assays Cancer science Medium 38403332
2025 Nuclear factor I A (NFIA) directly binds two motifs in the SMC4 promoter (-1379 bp and -354 bp), driving SMC4 transcription, as validated by dual-luciferase and ChIP assays. SMC4 in turn promotes G1/S cell cycle transition, activates TGF-β/SMAD signaling (upregulating p-SMAD2/3, N-cadherin, SNAI1, ZEB1) to enhance metastasis, and facilitates aerobic glycolysis by upregulating LDHA. ChIP assay, dual-luciferase reporter, ECAR/OCR metabolic assays, rescue experiments, xenograft and tail-vein metastasis mouse models, cell cycle analysis Frontiers in oncology Medium 40933894
2025 SMC4 is identified as a potential substrate of Polo-like kinase 1 (Plk1) in human cells; Plk1 activity is required for chromosome condensation and global chromosomal accessibility during mitosis, and SMC4 is proposed as a downstream effector in this regulatory mechanism. 5-ethynyl uridine nascent RNA labeling, Plk1 inhibition, ATAC-seq for chromatin accessibility, substrate identification by mass spectrometry bioRxivpreprint Low
2025 Solution AFM imaging of yeast condensin in various nucleotide states (AMP-PNP, ATPγS, ADP, no nucleotide) combined with coarse-grained MD simulations of Smc2/Smc4 heterodimer indicates that ATP binding at the head domains induces head engagement coupled to hinge opening, and that the N-terminal region of the Brn1 accessory subunit re-associates with the Smc2 head after ADP release. Solution high-speed AFM imaging, coarse-grained molecular dynamics simulation, flexible fitting to AFM images bioRxivpreprint Medium

Source papers

Stage 0 corpus · 64 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2016 ATPase-Modulated Stress Granules Contain a Diverse Proteome and Substructure. Cell 1233 26777405
2015 The BioPlex Network: A Systematic Exploration of the Human Interactome. Cell 1118 26186194
2017 Architecture of the human interactome defines protein communities and disease networks. Nature 1085 28514442
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2018 VIRMA mediates preferential m6A mRNA methylation in 3'UTR and near stop codon and associates with alternative polyadenylation. Cell discovery 829 29507755
2003 Complete sequencing and characterization of 21,243 full-length human cDNAs. Nature genetics 754 14702039
2007 Large-scale mapping of human protein-protein interactions by mass spectrometry. Molecular systems biology 733 17353931
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2012 A census of human soluble protein complexes. Cell 689 22939629
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2006 Wapl controls the dynamic association of cohesin with chromatin. Cell 498 17113138
1994 Oligo-capping: a simple method to replace the cap structure of eukaryotic mRNAs with oligoribonucleotides. Gene 492 8125298
2003 Differential contributions of condensin I and condensin II to mitotic chromosome architecture in vertebrate cells. Cell 460 14532007
2022 OpenCell: Endogenous tagging for the cartography of human cellular organization. Science (New York, N.Y.) 432 35271311
2015 Panorama of ancient metazoan macromolecular complexes. Nature 407 26344197
2004 14-3-3-affinity purification of over 200 human phosphoproteins reveals new links to regulation of cellular metabolism, proliferation and trafficking. The Biochemical journal 372 14744259
2015 Proteome-wide profiling of protein assemblies by cross-linking mass spectrometry. Nature methods 370 26414014
2006 Disrupted in Schizophrenia 1 Interactome: evidence for the close connectivity of risk genes and a potential synaptic basis for schizophrenia. Molecular psychiatry 345 17043677
2006 Phosphoproteome analysis of the human mitotic spindle. Proceedings of the National Academy of Sciences of the United States of America 281 16565220
2012 A high-throughput approach for measuring temporal changes in the interactome. Nature methods 273 22863883
2002 The ABCs of SMC proteins: two-armed ATPases for chromosome condensation, cohesion, and repair. Genes & development 265 11850403
2013 The functional interactome landscape of the human histone deacetylase family. Molecular systems biology 235 23752268
2017 Optimized fragmentation schemes and data analysis strategies for proteome-wide cross-link identification. Nature communications 221 28524877
1999 Fission yeast condensin complex: essential roles of non-SMC subunits for condensation and Cdc2 phosphorylation of Cut3/SMC4. Genes & development 221 10485849
2016 An organelle-specific protein landscape identifies novel diseases and molecular mechanisms. Nature communications 211 27173435
2015 ∆F508 CFTR interactome remodelling promotes rescue of cystic fibrosis. Nature 209 26618866
2013 PRP19 transforms into a sensor of RPA-ssDNA after DNA damage and drives ATR activation via a ubiquitin-mediated circuitry. Molecular cell 204 24332808
2018 An AP-MS- and BioID-compatible MAC-tag enables comprehensive mapping of protein interactions and subcellular localizations. Nature communications 201 29568061
2020 Systems analysis of RhoGEF and RhoGAP regulatory proteins reveals spatially organized RAC1 signalling from integrin adhesions. Nature cell biology 194 32203420
2017 The E3 ubiquitin ligase and RNA-binding protein ZNF598 orchestrates ribosome quality control of premature polyadenylated mRNAs. Nature communications 176 28685749
2001 A role for Drosophila SMC4 in the resolution of sister chromatids in mitosis. Current biology : CB 159 11267866
2000 A human condensin complex containing hCAP-C-hCAP-E and CNAP1, a homolog of Xenopus XCAP-D2, colocalizes with phosphorylated histone H3 during the early stage of mitotic chromosome condensation. Molecular and cellular biology 102 10958694
2023 The diapause-like colorectal cancer cells induced by SMC4 attenuation are characterized by low proliferation and chemotherapy insensitivity. Cell metabolism 78 37543034
2016 Condensin Smc2-Smc4 Dimers Are Flexible and Dynamic. Cell reports 70 26904946
2017 Overexpression of SMC4 activates TGFβ/Smad signaling and promotes aggressive phenotype in glioma cells. Oncogenesis 61 28287612
2014 A novel miR-219-SMC4-JAK2/Stat3 regulatory pathway in human hepatocellular carcinoma. Journal of experimental & clinical cancer research : CR 61 24980149
2016 SMC4, which is essentially involved in lung development, is associated with lung adenocarcinoma progression. Scientific reports 52 27687868
2015 Three-dimensional topology of the SMC2/SMC4 subcomplex from chicken condensin I revealed by cross-linking and molecular modelling. Open biology 45 25716199
2020 Direct DNA crosslinking with CAP-C uncovers transcription-dependent chromatin organization at high resolution. Nature biotechnology 37 32839564
2020 Plasmodium Condensin Core Subunits SMC2/SMC4 Mediate Atypical Mitosis and Are Essential for Parasite Proliferation and Transmission. Cell reports 29 32049018
2018 Condensin Smc4 promotes inflammatory innate immune response by epigenetically enhancing NEMO transcription. Journal of autoimmunity 25 29803706
2020 The Long Non-Coding RNA-RoR Promotes the Tumorigenesis of Human Colorectal Cancer by Targeting miR-6833-3p Through SMC4. OncoTargets and therapy 24 32273727
2017 Mutation of Arabidopsis SMC4 identifies condensin as a corepressor of pericentromeric transposons and conditionally expressed genes. Genes & development 24 28882854
2006 Disruption of the Arabidopsis SMC4 gene, AtCAP-C, compromises gametogenesis and embryogenesis. Planta 22 16482433
2022 TREM-1 induces pyroptosis in cardiomyocytes by activating NLRP3 inflammasome through the SMC4/NEMO pathway. The FEBS journal 20 36181338
2019 HIF-1-miR-219-SMC4 Regulatory Pathway Promoting Proliferation and Migration of HCC under Hypoxic Condition. BioMed research international 20 31828143
2021 MiR-433-3p restrains the proliferation, migration and invasion of glioma cells via targeting SMC4. Brain research 17 34147470
2019 Cell cycle regulation of condensin Smc4. Oncotarget 16 30719224
2021 SMC4 knockdown inhibits malignant biological behaviors of endometrial cancer cells by regulation of FoxO1 activity. Archives of biochemistry and biophysics 12 34506757
2024 FoxO1 promotes ovarian cancer by increasing transcription and METTL14-mediated m6A modification of SMC4. Cancer science 11 38403332
2021 Changes of EGFR and SMC4 expressions in triple-negative breast cancer and their early diagnostic value. Gland surgery 10 33842255
2022 SMC4 enhances the chemoresistance of hepatoma cells by promoting autophagy. Annals of translational medicine 9 36660610
2020 The condensin subunits SMC2 and SMC4 interact for correct condensation and segregation of mitotic maize chromosomes. The Plant journal : for cell and molecular biology 9 31816133
2024 ZNF131 facilitates the growth of hepatocellular carcinoma by acting as a transcriptional activator of SMC4 expression. Biochemical and biophysical research communications 5 38241815
2025 SMC4 Promotes Prostate Cancer Cell Proliferation and Metastasis via the Rheb/mTOR Pathway. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 3 40278414
2024 STAP-2 facilitates insulin signaling through binding to CAP/c-Cbl and regulates adipocyte differentiation. Scientific reports 3 38461189
2025 SMC4 promotes immune evasion by inhibiting endogenous interferon signaling and upregulating PD-L1 expression in triple negative breast cancer. International immunopharmacology 2 40976049
2025 USP39/SMC4 promotes hepatoma cell proliferation and 5-FU resistance. Scientific reports 1 40087331
2011 [Analysis for susceptibility of breast cancer due to gene SMC4L1 based on a multi-criteria evaluation model]. Sheng wu yi xue gong cheng xue za zhi = Journal of biomedical engineering = Shengwu yixue gongchengxue zazhi 1 21774228
2026 Superenhancer-driven SMC4 promotes myeloma growth by epigenetically enhancing IFI16-dependent STING signaling. Apoptosis : an international journal on programmed cell death 0 41793503
2026 SMC4/SMAD3/NF-κB axis drives cervical cancer progression and radioresistance via DNA damage repair and immune modulation. Journal of translational medicine 0 41803949
2025 RETRACTION: HIF-1-miR-219-SMC4 Regulatory Pathway Promoting Proliferation and Migration of HCC under Hypoxic Condition. BioMed research international 0 40538835
2025 NFIA-dependent upregulation of SMC4 promotes metastasis and metabolic reprogramming in glioma. Frontiers in oncology 0 40933894